The study of non-human primate thanatology has expanded dramatically in recent years as scientists seek to understand the evolutionary roots of human death concepts and practices. However, observations of how conspecifics respond to dead individuals are rare and highly variable. Mothers of several species of primate have been reported to carry and continue to interact with dead infants. Such interactions have been proposed to be related to maternal condition, attachment, environmental conditions or reflect a lack of awareness that the infant has died. Here, we tested these hypotheses using a dataset of cases of infant corpse carrying by chimpanzees in Gombe National Park, Tanzania ( n = 33), the largest dataset of such cases in chimpanzees. We found that mothers carried infant corpses at high rates, despite behavioural evidence that they recognize that death has occurred. Median duration of carriage was 1.83 days (interquartile range = 1.03–3.59). Using an information theoretic approach, we found no support for any of the leading hypotheses for duration of continued carriage. We interpret these data in the context of recent discussions regarding what non-human primates understand about death.

In many mammals, early social experience is critical to developing species-appropriate adult behaviors. Although mother-infant interactions play an undeniably significant role in social development, other individuals in the social milieu may also influence infant outcomes. Additionally, the social skills necessary for adult success may differ between the sexes. In chimpanzees (Pan troglodytes), adult males are more gregarious than females and rely on a suite of competitive and cooperative relationships to obtain access to females. In fission-fusion species, including humans and chimpanzees, subgroup composition is labile and individuals can vary the number of individuals with whom they associate. Thus, mothers in these species have a variety of social options. In this study, we investigated whether wild chimpanzee maternal subgrouping patterns differed based on infant sex. Our results show that mothers of sons were more gregarious than mothers of daughters; differences were especially pronounced during the first 6 mo of life, when infant behavior is unlikely to influence maternal subgrouping. Furthermore, mothers with sons spent significantly more time in parties containing males during the first 6 mo. These early differences foreshadow the well-documented sex differences in adult social behavior, and maternal gregariousness may provide sons with important observational learning experiences and social exposure early in life. The presence of these patterns in chimpanzees raises questions concerning the evolutionary history of differential social exposure and its role in shaping sex-typical behavior in humans.infant socialization | maternal behavior | chimpanzees | fission-fusion species E arly socialization is critical to developing social competency later in life. In mammals, mothers have enormous influence on their offspring's early social experience, with implications for adult social behavior. In humans, the relative contribution of parental and others' social influence on the development of sextypical behavior receives considerable attention and is often debated (1-4). Comparative research provides insight into the origins and development of sex differences in the absence of human cultural sex socialization. Decades of research in rodent and primate models have demonstrated that social deprivation curtails the development of species-appropriate behaviors (5-8) and cognition (9, 10). Primate mothers are critical for the normal social development of their infants (11-13), with classic studies demonstrating that maternal deprivation is associated with intense anxiety (14), inappropriate aggression (15), and an inability to form social relationships (16). The mother-infant relationship is therefore critical to proper social development; however, research also demonstrates the importance of the larger social milieu (17)(18)(19)(20). For example, a recent study in mice found that early interactions with mothers and peers independently shape adult behavior (21). Likewise, some negative impacts of maternal deprivati...

Early life experiences are known to influence hypothalamic-pituitary-adrenal (HPA) axis development, which can impact health outcomes through the individual's ability to mount appropriate physiological reactions to stressors. In primates, these early experiences are most often mediated through the mother and can include the physiological environment experienced during gestation. Here, we investigate stress physiology of dependent offspring in wild chimpanzees for the first time and examine whether differences in maternal stress physiology are related to differences in offspring stress physiology. Specifically, we explore the relationship between maternal rank and maternal fecal glucocorticoid metabolite (FGM) concentration during pregnancy and early lactation (first 6 months post-partum) and examine whether differences based on maternal rank are associated with dependent offspring FGM concentrations. We found that low-ranking females exhibited significantly higher FGM concentrations during pregnancy than during the first 6 months of lactation. Furthermore, during pregnancy, low-ranking females experienced significantly higher FGM concentrations than high-ranking females. As for dependent offspring, we found that male offspring of low-ranking mothers experienced stronger decreases in FGM concentrations as they aged compared to males with high-ranking mothers or their dependent female counterparts. Together, these results suggest that maternal rank and FGM concentrations experienced during gestation are related to offspring stress physiology and that this relationship is particularly pronounced in males compared to females. Importantly, this study provides the first evidence for maternal effects on the development of offspring HPA function in wild chimpanzees, which likely relates to subsequent health and fitness outcomes. Am. J. Primatol. 80:e22525, 2018. © 2016 Wiley Periodicals, Inc.

Objectives The prolonged juvenile period exhibited by primates is an evolutionary conundrum. Here we examine wild chimpanzee feeding development in the context of two hypotheses regarding prolonged development in primates: the needing‐to‐learn hypothesis and the expensive brain hypothesis. Material and Methods We studied wild chimpanzee (Pan troglodytes schweinfurthii) offspring at Gombe National Park, Tanzania. We analyzed 41 years of observational behavioral data collected between 1975 and 2016 from 81 offspring. We characterized feeding development in the first 10 years of life via four different measures: (1) proportion of observation time spent feeding; (2) diet composition; (3) diet breadth; and (4) diet maturity as measured by similarity to maternal diet. We used mixed effects models to examine changes with age and by sex, while controlling for season. Results Feeding time, diet breadth, and diet maturity exhibited the most substantial increases with age in the first 6 years, with no significant change thereafter. Males and females showed different patterns of change in diet breadth by age, but did not differ by age 10. Diet composition did not change significantly with age and did not differ by sex. Discussion We found that chimpanzee offspring attained adult‐like feeding behaviors between 4 and 6 years of age, concomitant with the completion of weaning. Thus, our data do not support the needing‐to‐learn feeding skills hypothesis of a prolonged juvenile period, but additional data are needed to evaluate how and when adolescent chimpanzees are able to make foraging decisions independent of their mothers. Existing data on growth provides support for the expensive brain hypothesis, however, these hypotheses are not necessarily mutually exclusive. As more studies across taxa accumulate sufficient datasets on a range of developmental metrics, we will be able to achieve a more robust understanding of prolonged development in primates.

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